Potential of high-density pheromone-releasing microtraps for control of codling moth Cydia pomonella

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1 Physiological Entomology (2012) 37, OI: /j x Potential of high-density pheromone-releasing microtraps for control of codling moth ydia pomonella and obliquebanded leafroller horistoneura rosaceana MIHEL. REINKE, JMES R. MILLER and LRRY J. GUT epartment of Entomology, Michigan State University, East Lansing, Michigan, U.S.. bstract. Recent large-cage studies with codling moth ydia pomonella (L.) reveal that the removal of moths from an apple orchard using pheromone-releasing traps is more effective at reducing capture in a central monitoring trap than is a mating disruption protocol without kill/capture. The present study uses open orchard 0.2-ha plots comparing a high-density trapping scenario with mating disruption to confirm those results. Two tortricid moth pests of tree fruit are studied: codling moth and obliquebanded leafroller horistoneura rosaceana (Harris). odling moth treatments include Isomate M FLEX (ShinEtsu Ltd, Japan), nonsticky traps baited with Trécé M lures (Trécé, Inc., dair, Oklahoma), and sticky traps baited with Trécé M lures, all at equal application rates of 500 dispensers ha 1, as well as a no pheromone control. These microtraps are of a novel design, small and easy to apply, and potentially inexpensive to produce. Mating disruption using Isomate M FLEX and nonsticky traps reduces codling moth capture in standard monitoring traps by 58% and 71%, respectively. The attract-and-remove treatment with sticky traps reduces capture by 92%. Obliquebanded leafroller treatments include Isomate OLR/PLR Plus and Pherocon II microtraps baited with Trécé OLR lures, both applied at 500 dispensers ha 1, as well as a no pheromone control. Mating disruption reduces capture in monitoring traps by 69%. The attract-and-remove treatment reduces capture by 85%. oth studies suggest that an attract-and-remove approach has the potential to provide superior control of moth populations compared with that achieved by mating disruption operating by competitive attraction. Key words. ttract-and-remove, mass trapping, mating disruption, microtraps. Introduction Semiochemically-mediated mass trapping as a control tactic has been of long interest for various lepidopteran species in agricultural systems. Roelofs et al. (1970) report that the reduction of fruit damage by moderate populations of redbanded leafroller rgyrotaenia velutinana is possible using monitoring traps ha 1 in apple orchards. Similarly, redbanded leafroller and grape berry moth Endopiza vitana orrespondence: Michael. Reinke, epartment of Entomology, Michigan State University, 106 IPS, East Lansing, Michigan 48911, U.S.. Tel.: ; reinkem3@msu.edu damage is reduced to moderate levels in grape vineyards using monitoring traps ha 1 (Taschenberg et al., 1974). oncerns over substantial costs of mass trapping systems (Huber et al., 1979; El-Sayed et al., 2006) and reduced individual trap effectiveness as a result of overlapping pheromone plumes (Madsen et al., 1976) have prompted tests with reduced trap densities. Willson & Trammel (1980) report using traps ha 1 in an unsuccessful attempt to control various tortricid species in apple orchards. Their study shows that damage from codling moth, in particular, can be reduced compared with the control, although it is still above acceptable levels for commercial practice. Madsen & arty (1979) also demonstrate the potential for codling moth control using traps at 10 and 36 ha 1, although the lack of replication and control Physiological Entomology 2012 The Royal Entomological Society 53

2 54 M.. Reinke et al. treatments precludes firm conclusions. Leskey et al. (2009) document low efficacy at trap densities of 5 and 20 traps ha 1 against dogwood borer Synanthedon scitula in apple orchards. Other recent studies with higher trap densities fare better. Zhang et al. (2002) provide control of the hinese tortrix ydia trasias in urban street plantings of hinese scholar-trees when traps are deployed at densities equivalent to 110 ha 1.ork et al. (2005) show that, in combination with other integrated pest management tactics, the control of a pyralid Leucinodes orbonali can be achieved using 100 traps ha 1. Eggplant production increases significantly compared with the standard, intense chemical programme. The consensus from these studies (El-Sayed et al., 2006) is that mass trapping has several hurdles to overcome before it can become a viable economic control tactic: the pheromone must be highly attractive and capable of drawing targets from long distances, yet able to bring them close to the source; the trap must be a design that is efficient at luring in and trapping the target species; the trap must be capable of capturing many individuals; and the traps must be inexpensive to allow many to be deployed per ha. Miller et al. (2006a, b) report that competitive attraction, or false plume following, is the dominant mechanism of semiochemical control among tortricid Lepidoptera. Under this mechanism, pheromone point sources directly compete with calling females. Over successive evenings, males of the targeted species will continue to orient towards individual pheromone plumes. To achieve control through competitive attraction, traps and dispensers must considerably outnumber females. This precludes the use of low numbers of dispensers or traps for controlling all but the lowest of population levels. In the present study, a high-density trapping regime is compared with mating disruption of codling moth ydia pomonella (L.) and obliquebanded leafroller horistoneura rosaceana (Harris). lthough the first of the four limitations to an effective attract-and-remove strategy listed above can be overcome by the use of commercially available lures, the others need to be addressed. In the present study, trap densities equivalent to standard hand-applied mating disruption dispensers are employed to test the hypothesis that attraction combined with male moth removal is superior to mating disruption. patent-pending, novel trap design is deployed against codling moth. It is suggested that this microtrap has the capacity to accomplish the attract-and-remove scenario without the compromises inherent with the large, complex and high-maintenance traps typically used for monitoring. small, commercially available, microtrap is used in the obliquebanded leafroller study. Materials and methods Orchards Research was performed at two Michigan State University research stations: larksville Horticultural Experiment Station and Trevor Nichols Research enter, located in Ionia and llegan counties, respectively. Two replicates were performed at larksville where treatments were placed in isolated 0.2-ha apple orchard plots. Each plot measured m and was surrounded by open grass or a border row of mature poplar trees. Each plot received a single treatment. Treatments were deployed in a randomized complete block design. ll orchard plots were trellis-planted at 1750 trees ha 1 and maintained as organic orchards but with no insect sprays. The two remaining replicates were placed in two 1.8-ha apple orchard blocks at Trevor Nichols Research enter. Orchards were divided into 0.2-ha plots measuring m set up diagonally from each other with a minimum of 15 m between treatments at the corners. Each 0.2-ha plot received one treatment. Treatments were deployed in a complete block design with randomization restriction. To prevent pheromone spread between treatment plots, treatments were deployed on a west east orientation from lowest total pheromone concentration to highest as an adjustment for the westernly daytime prevailing winds. ll plots contained freestanding apple trees planted at 500 trees ha 1. ll plots were maintained with regular horticultural practices but with no insecticide sprays. odling moth trapping Treatments included: (i) Isomate M Flex (ShinEtsu Ltd, Japan) applied evenly at 500 ha 1 (100 per plot); (ii) microtraps of a novel design described below without a sticky internal surface, each baited with one Trécé codling moth rubber septum (M L2; Trécé, Inc., dair, Oklahoma) lure, and applied evenly at 500 ha 1 (100 per plot); (iii) microtraps with a sticky internal surface, each baited with a rubber septum lure, and applied evenly at 500 ha 1 (100 per plot); and (iv) an untreated control plot. The nonsticky microtrap treatment was included to quantify any possible disrupting effect on moth movement. oing so permitted an assessment of the supplementary effect of removing moths using the sticky traps. Each plot received two monitoring traps (orange Trécé Pherocon VI delta traps, with a Trécé M L2 monitoring lure) placed in the southwestern and northeastern corners of the plot at least 15 m from the block perimeter. Treatments were applied on 7 May 2010 and were maintained until after the end of the second codling moth generation on 8 September The lures in the microtraps of treatment (iii) were changed between codling moth generations on 29 July For first-generation moths, Trécé M L2 monitoring lures were used. Low captures in the microtraps in first flight led to the suggestion that L2 lures release at an above-optimal rate to lure codling moth males into the microtraps. To test this hypothesis, all microtrap lures were changed to grey septum 0.1-mg lures produced by Trécé for the second codling moth generation. ll traps were checked weekly. Moths in monitoring traps were counted and removed. Moths captured in the microtraps were counted but were not removed and were designated as being caught on the bottom, side or top surfaces. Each microtrap was given a unique designation permitting the mapping of moth population differences within each plot. ll damaged or saturated traps and/or liners were replaced.

3 High-density pheromone-releasing microtraps 55 Microtrap design The microtrap is a 4-cm cube. The four sides and top have one hole (diameter 12 mm) in the centre of each surface. The bottom has two holes (diameter 5 mm) placed equidistant from each other and the sides of the microtrap. The purpose of the smaller holes on the bottom is to reduce the likelihood of the rubber septum and/or the captured moths from falling out of the trap. This microtrap design captures codling moths effectively in a flight tunnel (M.. Reinke, unpublished observations). Traps were constructed from modified lpha Scents (Portland, Oregon) plastic delta sticky inserts. clip made of bent steel wire was glued to one side of the cube for attaching microtraps to trees. Obliquebanded leafroller trapping Treatments included: (i) Isomate OLR/PLR Plus (ShinEtsu Ltd) applied evenly at 500 ha 1 (100 per plot); (ii) Trécé Pherocon II traps each baited with one Trécé OLR monitoring lure and applied evenly at 500 ha 1 (100 per plot); and (iii) an untreated control. The Trécé Pherocon II traps were the smallest available traps for attract-and-remove of obliquebanded leafroller as a result of the yet unproven efficacy of the cubical microtrap at capturing this species. Two orange Trécé Pherocon VI delta traps, with a Trécé OLR rubber septum monitoring lure, were placed in the southeastern and northwestern corners of the plot at least 15 m from the block perimeter. Treatments were applied on 1 June 2010 and were maintained until after the end of the second obliquebanded leafroller flight on 8 September ll traps were checked weekly. Moths in monitoring traps were counted and removed. Moths captured in all other traps were counted but were not removed. Each Trécé Pherocon II trap was given a unique designation and individually maintained to facilitate mapping of moth population differences within each plot. ll damaged or saturated traps and/or liners were replaced. Shoot damage assessment was performed July Ten shoots were checked on each of 30 trees per plot. Each shoot tip was inspected for the presence of larvae and/or feeding damage. Statistical analysis For orientation disruption and trapping studies, weekly moth capture in the two traps in each plot was summed. Weekly data were log transformed [ln (x + 1)] (which normalized the distributions and homogenized variance) and then subjected to analysis of variance (anova). Trapping treatment trap captures were summed by generation for individual traps, and then log transformed [ln (x + 1)] and subjected to anova. Shoot injury data were log transformed [ln (x)] before anova. ifferences in pairs of means were separated using the least significant difference test (SS Institute, 2000). In all cases, α < 0.05 was considered statistically significant. Results odling moth trapping For the first generation, total moth captures for the Isomate, nonsticky microtrap and sticky microtrap treatments were significantly lower than for the untreated control (P < 0.001, P<0.001 and P<0.001, respectively) (Fig. 1). No significant difference was observed between the Isomate and nonsticky trap treatments (P = 0.089). apture in monitoring traps was significantly lower in the sticky microtrap plots than both the Isomate and nonsticky microtrap treatments (P < and P<0.001, respectively) (Fig. 1). Weekly captures in the trapping treatment were summed for each trap in the first codling moth generation. These counts were mapped using a surface map (Fig. 2). For the second generation, total moth captures in monitoring traps for the Isomate, nonsticky microtrap and sticky microtrap treatments were significantly lower than for the untreated control (P <0.001, P<0.001 and P<0.001, respectively) (Fig. 1). No significant difference was observed between the Isomate and nonsticky trap treatments (P = 0.488). apture in monitoring traps was, however, significantly lower in the sticky microtrap plots than the Isomate plots (P = 0.023) but not the nonsticky microtrap plots (P = 0.108) (Fig. 1). Weekly captures in the trapping treatment were summed for each trap in the second codling moth generation. These counts were mapped using a surface map (Fig. 3). Microtrap design ll interior surfaces of the microtrap were sticky. However, only the sides and bottom effectively captured high numbers of codling moths. Of the 644 males captured in the microtraps during the first generation, 60% were captured on the bottom interior surface (Table 1). The remaining 40% were captured on the four sides. uring the second generation, total capture increased significantly (P <0.001) to 1308; the bottom captured 53% of moths. The remaining 46% and 1% were captured on the sides and top, respectively. verage catch in monitoring traps ±SE First generation Second generation ontrol Isomate N.S. Trap Sticky Trap Fig. 1. apture ofcodling moth ydia pomonella males in monitoring traps. Mean catch per plot is used with catches summed for both monitoring traps in each plot, for each generation. N.S. Trap, nonsticky trap.

4 56 M.. Reinke et al. Table 1. Total number of codling moth ydia pomonella males captured in microtraps, including a breakdown of catches on sections of traps. apture (% grand total) Generation ottom Sides Top Grand total One 386 (60%) 258 (40%) Two 689 (53%) 601 (46%) 18 (1%) 1308 Total 1075 (55%) 859 (44%) 18 (1%) Fig. 2. Surface map showing cumulative number of codling moth ydia pomonella males captured in each microtrap in the trapping treatment during the first generation. olours designate cumulative moth counts in each trap. Large surfaces with the same colour are indicative of multiple traps with identical counts. Parts () to () indicate the individual replicates shown in Figs 8 and 9. trapped plots more than the Isomate plots (P = 0.049). Weekly captures in the trapping treatment were summed for each trap in the first obliquebanded leafroller generation. These counts were mapped using a surface map (Fig. 5). For the second generation, total moth captures in the monitoring traps for the Isomate and trapping treatments were reduced by 81% and 94%, respectively, compared with the untreated control (Fig. 4). These reductions were both significant (P <0.001 and P<0.001, respectively). aptures in monitoring traps were reduced significantly in the trapped plots more than in the Isomate plots (P = 0.007). Weekly captures in the trapping treatment were summed for each trap in the second obliquebanded leafroller generation. These counts were mapped using a surface map (Fig. 6). Mid-season shoot damage was significantly reduced in the Isomate mating disruption and high-density trapping treatments (P <0.024 and P<0.003, respectively) (Fig. 7). No significant difference was found between these pheromone treatments (P = 0.099) iscussion El-Sayed et al. (2006) suggest that traps at high density would operate more as mating disruption dispensers than as traps. This implies that mass trapping cannot out-perform mating disruption. However, computer simulations comparing mating disruption with mass trapping (yers, 2007) and research on mating disruption mechanisms conducted in large field cages (Miller et al., 2010) do not support this proposition. In the latter study, known numbers of codling moths are released Fig. 3. Surface map showing cumulative number of codling moth ydia pomonella males captured in each microtrap in the trapping treatment during the second generation. olours designate cumulative moth counts in each trap. Large surfaces with the same colour are indicative of multiple traps with identical counts. Parts () to () indicate the individual replicates shown in Figs 8 and 9. Obliquebanded leafroller trapping For the first generation, moth captures in the Isomate and trapping treatments were reduced by 57% and 76%, respectively, compared with the untreated control (Fig. 4). These reductions were both significant (P <0.001 and P< 0.001, respectively). aptures were significantly reduced in the verage catch in monitoring traps ±SE First generation Second generation ontrol Isomate Trap out Fig. 4. apture of obliquebanded leafroller horistoneura rosaceana males in monitoring traps. verage catch per plot is used with catches summed for both monitoring traps in each plot for each generation.

5 High-density pheromone-releasing microtraps 57 verage percentage shoot damage ±SE ontrol Isomate Trap out Fig. 7. Percentage of mid-season shoots damaged by obliquebanded leafroller larvae horistoneura rosaceana Fig. 5. Surface map showing cumulative number of obliquebanded leafroller horistoneura rosaceana males captured in each trap in the trapping treatment during the first generation. olours designate cumulative moth counts in each trap. Large surfaces with the same colour are indicative of multiple traps with identical counts. Parts () to () indicate the individual replicates shown in Figs 8 and Fig. 6. Surface map showing cumulative number of obliquebanded leafroller horistoneura rosaceana males captured in each trap in the trapping treatment during second generation. olours designate cumulative moth counts in each trap. Large surfaces with the same colour are indicative of multiple traps with identical counts. Parts () to () indicate the individual replicates shown in Figs 8 and 9. into large field cages with known dispenser and trap densities in the range ha 1. conclusion for codling moth is that standard hand-applied mating disruption is more effective than lures placed in linerless disrupting traps with respect to reducing the capture of males in monitoring traps. Without liners, the lure/trap combination is operating as a low-releasing dispenser. Once liners are added to the traps, however, there is a supplementary decrease in the capture of males compared with the standard disruption. t the highest density of sticky traps, capture in the central monitoring trap is reduced by fourfold (75%) compared with the hand-applied dispensers and by nine-fold (89%) compared with the nonsticky low-releasing dispensers (Miller et al., 2010). Miller et al. (2010) deduce that high rates of pheromone released from the hand-applied dispensers deactivate attracted males, whereas low-releasing dispensers do not. This temporary deactivation eliminates further orientation by individuals on a given evening but allows orientations during the next flight period. The addition of a capturing medium results in permanent deactivation. The present codling moth open-field study confirms and extends the large-cage work. aptures in the Isomate and nonsticky microtrap plots are reduced by 58% and 71%, respectively, compared with the untreated plot (Fig. 1). ecause nonsticky microtraps depressed catch in the monitoring traps to levels equal to Isomate dispensers, it is tentatively concluded that males are retained for most of each activity period. Laboratory flight-tunnel studies indicate this trap design is capable of retaining male codling moths for longer than a large plastic delta trap (M.. Reinke, unpublished observations). This effect may be a result of the design, where, in the case of the microtrap, the male has a flat surface to alight upon at the source of the pheromone plume but a small orifice to enter and exit the trap. Once the moth crawls into the nonsticky trap, the small orifices make exit difficult. Retaining the male codling moths inside the nonsticky traps for an extended time would equate to nightly deactivation. This condition would, however, be temporary, just as it is for the hand-applied rope dispensers. s expected, the addition of a capturing medium to the inside of the trap enhances the reduction of capture in the monitoring traps compared with all mating disruption treatments. aptures of codling moth males in the sticky microtrap plots are reduced by 92% compared with the control and by 80% and 72% compared with the hand-applied and nonsticky microtrap treatments, respectively. apture of obliquebanded leafroller in the trapping treatment is reduced by 85% compared with the control and by 51% compared with the mating disruption treatment. oth studies confirm that the attract-and-remove tactic can provide a superior control of codling moth and obliquebanded leafroller populations in apple orchards.

6 58 M.. Reinke et al. annual annual annual Fig. 8. Google Earth map showing placement of trapping treatment replicates for both codling moth ydia pomonella and obliquebanded leafroller horistoneura rosaceana at larksville Horticulture Experiment Station, larksville, Michigan. Uppercase letters and designate individual replicates. home lawns pears cherries Fig. 9. Google Earth map showing placement of trapping treatment replicates for both codling moth ydia pomonella and obliquebanded leafroller horistoneura rosaceana at Trevor Nichols Research enter, Fennville, Michigan. Uppercase letters and designate individual replicates. The microtrap is effective at capturing codling moth males. The higher captures during the second generation can be attributed to the lower release rate of the 0.1-mg lure (Figs 2 and 3 and Table 1). The M L2 lure used during the first generation probably releases at a rate not conducive to optimal male entry into the trap. If the 0.1-mg lure had been used over the full season, even more male codling moths would have been captured during the first generation. The vertical sides and horizontal bottom of the microtrap are equally important in capturing the high numbers of moths (Table 1). The top interior sticky surface is ineffective at capturing moths but could cause them to fall into the trap. dditional work is necessary to determine whether a sticky top contributes to overall trap efficiency. Midseason shoot damage assessment confirms the monitoring trap counts indicating that both mating disruption and highdensity trapping are effective at diminishing obliquebanded leafroller populations. odling moth fruit damage could not be assessed as a result of infestations of apple flea weevil that lead to a negligible fruit set. Fruit counts in all plots at larksville Horticultural Experiment Station are reduced to levels that are too low for differentiation between mating disruption and trap-out. dditional investigations into codling moth control using high trap densities and an attract-and-remove strategy are needed. eploying traps at high densities has the added benefit of reducing the burden of each individual trap from capturing all nearby moths. t a density of 500 traps ha 1, the largest distance between traps is 5 m. s such, plume overlap is likely to occur (Murlis et al., 2000; yers, 2007; Yamanaka, 2007). This suggests that, under a competitive attraction scenario, a male moth has the potential to encounter the plumes of multiple traps. t high moth densities, where seasonal capture would be approximately 2000 moths ha 1, as indicated in the present study as well as in previous studies (Roelofs et al., 1970; Madsen & arty, 1979), mass trapping with low trap densities is likely to result in trap saturation. With the trap densities reported in the present study, those catches would be shared by a larger number of traps. If one trap begins to saturate, surrounding traps would be capable of capturing those moths escaping the saturating microtrap. Trap saturation at these trap densities is doubtful, however, when capturing codling moth or obliquebanded leafroller males. In the unlikely event that a trap did saturate, it would still have a substantial effect as a mating disruption dispenser, as indicated by the activity of the nonsticky microtrap treatment. Figures 2, 3, 5 and 6 provide a visual record of population density on a fine scale. Hotspots are easily discernible by islands of contrasting colour against the blue background. These population centres envelop two to four traps. It appears that the hotspots are confined to no more than two to four trees in low-density apple plantings, and five or six trees in one or two rows in a trellis designed orchard. y placing traps at high densities, it is possible to have at least one trap in each of these high population centres, facilitating earlier capture and removal from the population. In addition to revealing hotspots, the surface maps assist with the visualization of population dynamics on a nearindividual-tree scale. odling moth males were captured throughout the blocks, although there is a marked edge effect in all four replicates (Figs 2, 3). In three of the four replicates, there is a marked increase in moth capture during the second generation. This is a result of multiple factors. First, the lures used in the microtraps were changed between generations to one with a release rate probably more conducive to moth entry into the trap, thus increasing the capture of a larger proportion of responding males. This is supported by comparing the generation-specific captures in the monitoring traps. lthough capture in the microtraps increases significantly during the

7 High-density pheromone-releasing microtraps 59 second generation, capture in the monitoring traps decreases (Fig. 1). Second, historically, both research stations have high codling moth populations during the second generation in unmanaged blocks. Replicate at larksville (Fig. 8) and both replicates at Trevor Nichols Research Station (Fig. 9) are in close proximity to apple orchard blocks. In all three cases, the largest moth captures are along the edges adjacent to these other apple blocks. These conditions facilitate immigration pressure. Even at the high trap densities tested, the pressure is too high to prevent immigration into the interior of the blocks. lternatively, replicate at larksville is isolated from other apple orchard blocks (Fig. 8). The nearest untreated apple block is 300 m to the southwest, on the other side of a stand of mature hardwood trees. This reduces immigration pressure. aptures in the microtraps in replicate decrease in the second generation, even with the higher trap efficiency as a result of new lures. Spatial maps of obliquebanded leafroller captures suggest a similar population structure (Figs 5 and 6). The use of the same lure type for both generations allows the captures from both generations to be directly compared. Most catches during the first generation occur at the perimeter of each plot. Male capture in Replicate increases slightly during the second generation. aptures in the remaining three replicates markedly decrease during the second generation. In all replicates, all traps with cumulative captures of three or more moths per trap during the second generation are within 10 m of the block perimeter. These maps indicate that obliquebanded leafroller populations are successfully controlled using an attract-and-remove programme. Obliquebanded leafroller captures appear to be primarily the result of immigration into the experimental plot. lthough the novel microtrap shows promise for use in an attract-and-remove regime for the codling moth, further development is required. The high capture rates reported in the present study indicate that the microtrap has the potential to efficiently capture codling moth males. Future work aims to test its efficacy against other lepidopteran pest species in various agricultural systems. The microtrap is small and constructed of inexpensive materials, although the development of a cost-effective assembly process is required before microtraps can be considered economically viable. cknowledgements We thank P. McGhee and J. Huang for their assistance in experimental design; L. Jagenow and M. Julian for helping check hundreds of traps weekly; and R. Isaacs and. Guyer for improving an earlier version of the manuscript. References yers, J.. 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